J. A. Bergen: Beniassian to Early Aptian Calcareous Nannofossils. .., p. 59-69.

Journal ofNannoplankton R~arch, 16, 2, 1994.

BERRIASIAN TO EARLY APTIAN CALCAREOUS NANNOFOSSILSFROM THE VOCONTIAN TROUGH (SE FRANCE) AND DEEP SEADRILLING SITE 534: NEW NANNOFOSSIL TAXA AND
A SUMMARY OF LOW-LATITUDE BIOSTRATIGRAPHIC EVENTSJames A. Bergen, Amoco Production Company, Houston , Texas
Abstract: Recent investigation of 17 sections in the Mesozoic Vocontian Trough (southeastern France) spanning theCallovian to Cenomanian has shown great potential for improving Tethyan nannofossil biostratigraphic resolution.
Both predefined zonal taxa and newly-recognized biostratigraphic events were calibrated to the Lower Cretaceousammonite biostratigraphy. Coeval strata drilled at Deep Sea Drilling Program Site .534 offered the chance to test theviabil ity of these new events and evaluate the calibration of the deep~ nannofossil biozonation. Reported here isa summary for the Berriasian to lower Aptian, with the most significant improvements being for the Valanginian andHauterivian. Ten ne w species and two new combinations are introduced.

Introduction
The thick basinal sequence of the Mesozoic Vocontian
Trough in southeastern France contains a fairly complete
record of nannofossil evolution and now provides an
excellent opportunity to calibrate the nannofossil record to
the Tethyan ammonite biostratigraphy. The Vocontian
Trough also contains several of the stage-stratotypes(Berriasian, Biuremian, and Aptian ; Valanginian
hypostratotype) and has been the subject of recent sequence stratigraphic work. The first published Lower
Cretaceous nannofossil biozonation by Thierstein ( 1971 ,1973) also was founded on sections in this basin. Bralower,
Monechi, and Thierstein ( 1989) improved biostratigraphic
resolution for the Tithonian to Berriasian and were able totie the nannofossil biostratigraphy to both the Tethyanammonite biostratigraphy and palaeomagnetic record.
Deep Sea Drilling Project Site 534 is an important reference section for M-sequence palaeomagnetic record. Roth(1983) published the original nannofossil biostratigraphyof this site, for which he assigned numbers his Lower
Cretaceous zones (NCl-10). Sissingh (1977, 1978) pu~
lished a numbered, zonal scheme for the Cretaceous (CCZones); Applegate and Bergen ( 1988) further refined thiszonal scheme based on Lower Cretaceous sections drilledon the Galicia Margin during O.D.P Leg 103.
This paper describes the new species found during
study of the Lower Cretaceous sections in the Vocontian
Trough and D.S.D.P Site 534. An outline of the sequenceof nannofossil events in common between these two
localities is provided and correlated to the ammonite
biostratigraphy of the French sections. Comparisons also
are made to the aforementioned nannofossil biozonations.
The taxonomic problems encountered are beyond thescope of the current paper and, hopefully, will be addressed in a future publication on the entire
Callovian-Cenomanian interval.

Methods
Research on the Vocontian Trough sections was completed during 1991 at Amoco Production Research in
Tulsa, Oklahoma. Two factors had a significant impact on
the results obtained during this study: centrifugation ofsamples and analysis time. These factors improved thecontinuity of stratigraphic ranges, especially for many of
the rare taxa.
Abbreviations used in this paper include: LM forlight microscope; SEM for scanning electron microscope;
LO for lowest occurrence; HO for highest occurrence; TR
for total range; Sa for samples; T for top; B for bottom; andSZ for Sub zone . Two rough estimates for thebiostratigraphy reliability of an individual species arecited: ( 1) percentage of samples in which the species wasfound throughout its entire documented stratigraphic
range; or (2) the number of samples in which the species
was found in either the top or bottom ten samples of its
documented stratigraphic range.

Acknowledgments
Special thanks to Luc Bulot for his interpretations of theammonite biostratigraphy. I also would like to thankAmoco Production Company for the opportunity to workon this research project, Eric de Kaenel for his help withthe specimen transfer (LM/SEM) documentation of specimens illustrated herein, and Tim Bralower for reference
samples from the BGS Speeton Borehole and DSDP
Site 534. Thanks once again to Jeremy Young, as well as
Paul Bown and David Rutledge for their helpful suggestions and reviews of this article.
Sections Considered
TheprimarysectionsforthecurrentpaperincludeD.S.D.P.
Site 534 (Biake-Bahama Plateau, NW Atlantic) and four
outcrops in SE France (see Text-Figure 1): (1) Berriasianto Valanginian at Angles; (2) Hauterivian at Vergons; (3)Barremian stratotype at Angles; and (4) the lower Aptian
stratotype at La Bedoule. Also referenced are the Speeton
section in Great Britain, the BGS Speeton Borehole, as
well as other sections in SE France (see Text-Figure 1):(A) Albian near Blieux; (B) upper Aptian at Angles; (C)the Berriasian stratotype; (D) Tithonian to lower Berriasiansections at Broyon and Le Pouzin (Route de Grads &Carriere de Anges); and (E) Upper Jurassic sections in SE
France at La Melle and Lavilledieu. Relevant nannofossil
publications include: Thierstein ( 1971 , 1973) on the
Barremian stratotype, Berrias, and Broyon; Bralower etal. (l989)onBerrias, Broyon, andD.S.D.P. Site534; Roth(1983) on D.S.D.P. Site 534; and the most recent paperon the Speeton section (Crux, 1989). The; BerriasianBarremian ammonite biostratigraphy is based on Bulot(1993 & 1994, pers. comm.) and the published frameworkof Hoedemaeker & Bulot (reporters) (1990) andHoedemaeker & Company (1993). The lower Aptian===========================59===========================

Journal ofNannoplanldon Research , 16, 2, 1994.

owns •
Sections •J. A. Bergen: &m.ssian to Early Aptian Calcareous Nannofossils. .. , p. 59-69.

0

=

~N

Te:rt-flcure I: Location of Vocontian Trough sections in southeast France. See text (under Sections Considered) for numbering and lettering ofsections.

ammonite zones were taken from Busnardo ( 1984) and aguide prepared for an Amoco field excursion (Busnardo etal., 1987).

Biostratigraphic Summary
A sequence of nannofossil events observed in the composite Vocontian Trough section and their correlation to theammonite biostratigraphy are summarized in the TextFigure 2. Only four of these events, which were very
distinct and reliable in the French sections, were observed
out of sequence at D.S.D.P. Site 534. Sample density(-1 Sa/5m) at Site 534 was a limiting factor (see Appendix
for list of samples). The thicker Vocontian sequence wasdensely sampled (1 Sa/2 m); facies and sampling density
were only problematic for the lower-middle Berriasian.
The following nannofossil events supplemented ammcr
nite correlations of the four primary Vocontian Troughsections (see Table in Appendix): (1) LO of Hayesites
irregularis for the Barremianand lower Aptian stratotypes;(2) HOofLithraphidites bolli i for the Barremian stratotype
and Vergons; (3) HO of Eiffellithus windii for Vergons
and Angles (Berriasian-Valanginian section).
Systematics: New Species and New Combinations
Assipetra terebrodentarius (Applegat~. Bralower,
Covington & Wise, 1987) Rutledge & Bergen,new combination
Remarks: Small specimens transitional between thisspecies andAssipetra infracretacea (Thierstein) occur inthe upper Hauterivian.
Jlange: LO is uppermost Hailterivian in: (1) DSDP Site
534 (100% Sa), Core 51-1 , 61-64cm; (2) SE France,
Angles, Barremian stratotype, Bed 70, angulicostataZone(76% Sa); (3) Speeton (approximate), Bed LB5 .

===========================60===========================

J. A. Bergen: Berri~ to Early ~an Calcareous Nannofossils. .., p. 59-69.

Calcicalathina, species A(Plate 1, figures lla, b)
Remarks: This large (9-1 0 J..lm), elliptical murolith is aform transitional between Rhagodiscus asper (Stradner,
1963) Reinhardt, 1967 and Calcicalathina oblongata
Thierstein, 1971 . It differs from the former species by its
large size and coarsely granular (more birefringent),
elevated central area. It differs from the latter species by
its wider rim (more optically distinct) and lower central
area. In cross-polarized light, Calcicalathina oblongata
displays an irregular longitudinal extinction line and its
poles uniform, bright birefringence (see Thierstein, 1971,
pl. 4, figs . 6-10 for comparison).
Range: earliest Berriasian to latest Aptian. DSDP Site
534 : LOin 90-3, 21-22cm. SE France: LOin grandis SZ(Route de Grads, Bed 27), 0.5m below grandis SZ (Broyon,
19.5m); HO at Angles, Bed 77, upper Clansayesian.

Diloma, species A(Plate 1, figures 12a, b)
Remarks: This medium-sized species of Diloma has aperforate central area spanned by an axial cross. Other
Diloma species ·with axial central structures are differentiated by the presence of diagonal struts in the central area
and/or inner rim margin. These accessory bars are restricted to the inner rim margin in Diloma prim.itiva(Worsley, 1971) and beautifully illustrated by Wind &tepek (1979; pl. 8, figs. 2, 3). However, such structures
are not mentioned in the original description of thisspecies and only barely visible in one (Worsley, 1971, pl.
1, fig. 2) of the three photographs of the holotype. The
original description of Diloma primitiva also mentions animperforate central area.
Range: Cosmopolitan, early Valanginian to early
Barremian. DSDP Site 534 : Sa 76-6, 94-95cm; SE France:
LO at Angles, Bed 289, carnpylotoxum Zone; HO at
Vergons, Bed 146, ligatus Zone; Speeton (approximate),
Beds D2 to LB5.

Diloma galiciense, new species(Plate l , figures l3a-c, l4a-c)

Journal ofNannoplanklon Research, 16,2, 1994.

tral area margin, giving the impression of a tricyclic rim
extinction pattern. A central cross spans the central
opening and its arms flare as they attach to the proximal
plate. The central cross is oriented 10-30 degrees off the
major ellipse axis and becomes brightly birefringent when
the coccolith is oriented 45 degrees to the polarizing
direction. A fibrous, distal projection is present at thecentre of the cross. Size of measured specimens: 7.0-9.0

J.lm.
Remarks: Diloma galiciense is distinguished from other
Diloma species by its diagonal central structure and fromother Neocomian muroliths by its tricyclic rim extinction
pattern.
Range: Cosmopolitan. DSDP Site 534 (58% Sa), lower touppermost Hauterivian, 63-2, 66-69cm to 52-1, 82-85cm;
SE France (Vergons, 25% Sa), Hauterivian, Bed 42(radiatus Zone) to Bed 89 (sayni Zone). North Sea(Rutledge, written comm., 1994), lower Hauterivian tolower Barremian.
Holotype: Applegate & Bergen (1988), plate 8, figs . 5, 6,
8; ODP Site 638B-25R-4, 48-49cm; upper Hauterivian.

Grantarhabdus quadratus (Worsley, 1971)
new combination(Plate 1, figure 10)1971 Watznaueria quadrata- Worsley, p. 1315, pi. 2, figs. 20-22.

Remarks: This species has a diago~, sigmoid crossstructure, small central area, and a thick proximal shield.
It is assigned to the genus Grantarhabdus based on its
central structure orientation, although its rim construction is more typical of the genus Helenea (junior synonym:
Microstaurus) .
Range: Cosmopolitan, earliest Berriasian to late
Hauterivian. DSDP Site 534 (88% Sa) : 91-2, 145-146cm
to 55-3, 75-78cm. SE France: LO at Le Pouzin, Bed 27,
grandis SZ; LO at Broyon, 19.5m, 0.5 m below grandis SZ;
HO at Vergons(87%Sa), Bed ll9, ligatusZone. Speeton(approximate): HO in Bed CS.

Hexalithus strictus, new species(Plate 1, figures la-b, 2a-b)

1988 Vekshinella angusta (Stover) Verbeek, 1977b. Applegate& Bergen,p. 318, pi. 8, figs. 5, 6, 8.
1991 Chiastozygus sp. Bralower, p. 430, fig. 6, nos. 31 ·32.

· Derivation of name: after the Galicia Bank, offPortugal.
Diagnosis: Eiffelithid exhibiting a bicyclic rim extinction
pattern (outer faint/inner bright) and having in its central
area an off-axis central cross attached to a proximal plateof faint birefringence.
Description: Medium to large, elliptical murolith. The
rim is bicyclic in distal view. The inner rim cycle isconstructed of blocky elements and is brightly birefringent.
The narrow outer rim cycle is faintly birefringent and its
elements show strong dextral imbrication. A faintly
birefringent proximal plate circumscribes the outer cen-

Derivation of name: Latin. strictus = straight.
Diagnosis: A species ofHexa/ithuswith triangular-shaped
elements (i.e . their peripheries are straight).
Description (LM only): Medium-sized, nannolith constructed of six, triangular, non-imbricated elements
separated by radial sutures. The individual element peripheries are straight, giving the nannolith a hexagonal,
peripheral outline. Size of observed specimens: 4.8-6.0

J.lm.

~,;

R~marks: Hexalithus noelae Loeblich & Tappan, 1966
has elements with lobate peripheral margins. Hexalithus

strictus has a younger appearance and extinction.

==========================61========================~

Journal ofNamoplankton Research , 16, 2, 1994.

J. A. Bergen: Berriassian to Early ~an Calcareous Nannofossils... , p. 59- 69.

~ONITE SE FranceiDSDP 534~ ZONFJSUBZONE

C'-lgrandis~ b hambrovi
matheroni~~ .S

Bf- 1st events Bf- 2nd events U!A-- E.noraliav-- N.ste1nmannll

angulicostata
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52-3
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A- R .tmgunu•~ R.angustiforata/C.mexicanalA-- Z.scutula~ ~ loryi- - Range difference 534/SE France~~<~~-~ :::>~E ~:g ·;;;~ ~ ·u
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coquandi"Prodesbaves!les"H.lrregularllsarasini
111.1raudi~~ R .achlyostaurion!':: feraud.tanus::1 sartoustana
L.salomonii
I'. vandcnbcck 1
FR.radiata::::::::::::::::::::::::::::.
............................
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ruCk estv-- E .hauterivianus
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~

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.
N.colomi CCI NCl• Applegate&Bergen (88)................., moutonianum/compressissima Zones

Tnt-Fipre l: Stages divisions and subdivisions and anunonite zones based on SE France; Berriasian to Barremian ammonite zones after Hoedemaeker& Bulot (1990) and Hoedemaeker & Company (1993) as communicated by Bulot (1993 , 1994); Aptian ammonite zones after Busnardo ( 1984) andBusnardo et al. ( 1987). The third column summarizes the sequence ofnannofossil events in SE France and correlation to ammonite zones; reliability indices(see methods) indicated are: ( 1) primary (1st), more than 90% Sa; and (2) secondary (2nd), more than 70% Sa. Correlation 5t-'i!annofossil events to DSDP
Site 534 is shown at the right side of column three and the Core-Sections are indicated in the fourth column. The nannofossil zones ofThierstein ( 1971,1973), Sissingh (1977, 1978), Roth (1978, 1983), Applegate & Bergen (1988) are illustrated in the fifth column relative to the sequence ofnannofossil
events; the slanted NC3/NC4 Zone boundary is due to variance in the HO ofT. verenae. The stratigraphic ranges of other important nannofossil species
are shown on the right side ofthe figure (relative to sequence ofn81Ul0fossil events); differences in tl. ir strati graphic ranges between SE France and DSDP
Site 534 are shown as dashed lines.

========================~62========================~

J. A. Bergen: Berriassian to Early Aptian Calcareous Namofossils. .. , p. 59-69.

Range: earliest Berriasian; DSDP Site 534 (55% Sa):
Cores 92-4, 147-148cm to 89-4, 35-36cm; SE France: Le
Pouzin-Route de Grads, Bed 25 <iacobi SZ) and Bed 27(grandis SZ); Broyon, 19.5m, (0.5m below grandis SZ).
Holotype: Plate 1, figs. la-b; DSDP Site 534, 91-4, 131l32cm; lower Berriasian.
Lucianorhabdus salomonii, new species(Plate 1, figures 17a-b, 18a-b)

Derivation of name: named in honour of Ralph A.
Salomon.
Diagnosis: A species of Lucianorhabdus with a massive,
truncated distal stem which displays a bright, longitudinalband down its central portion.
Description (LM only): Elliptical holococcolith having
a tall, broad distal stem which fills the entire central area.
The stem has a thin axial canal and a truncate distal
extremity. Its vertical outline is somewhat variable, including: (1) slight or no distal taper; and (2) a straight or
curved shape. In cross-polarized light, the stem displays
a characteristic, bright longitudinal band down its central
portion, whereas the remainder of stem is faintly
birefringent. The rim elevation is normal to high (height- 112 the coccolith width) and its wall diverges in the distal
direction. The narrow rim displays a first order whitebirefringence in both plan and lateral view. In plan view,
isolated rims without distal projections have their imperforate central areas divided into four quadrants by diagonal
extinction lines. Coccolith size: 4.5-6.0 !J.m; stem heightabout 10 !J.m.
Range: DSDP Site 534: Core 45-4, 10-l3cm, upper
Barremian; SE France (Angles): upper Barremian (Bed161, sartousiana Zone) to basal upper Aptian, Angles,
Bed 46.
Holotype: Plate 1, figs. 17a-b; Angles, upper Barremian,
Bed 161 (top sartousiana Zone).
Markalius vetulus, new specie~(Plate 1, figures 22a-b, 23a-b)

Derivation of name: Latin. vetulus (dim.)= somewhat
old.
Diagnosis: A Neocomian Markalius species having asteep, brightly birefringent tube cycle and shields thatexhibit a faint, smooth birefringence.
Description (LM only): Medium-sized, sub-circular tocircular placolith constructed of two shields, which areboth faintly birefringent and have indistinct sutures. The
proximal shield size is about 3/4 that of the distal shield.
Shield elements (-32 in number) are radial with slightcurvature near the outer periphery. A steep tube cycle
nearly encloses the central area (-113 coccolith diameter)
and exhibits a bright, 1st order white birefringence. Sharp
extinction lines cross the tube cycle, but do not extend onto
the shields. A very small central perforation is present.

Journal ofNannoplankton Research , 16, 2, 1994.

Observed specimens: 6.0 to 7.5 !J.m.
Remarks: Markalius inversus (Maastrichtian-Palaeogene)is very similar. However, its birefringent tube cycle is notsteep and its shield elements appear striated (i.e. individual elements are easily distinguished with the lightmicroscope).
Range: late early Valanginian; DSDP Site 534 (57% Sa) :Cores 79-2, 10-11cm to 76-6, 94-95cm; SE France (Angles, 80% Sa): Beds 278 to 302 (campylotoxum Zone).
Holotype: Plate 1, figs . 22a-b; Angles, Bed 278, upper
lower Valanginian.
Percivalia bulla/a, new species(Plate 1, figures 4a-b, 5a-b, 6a-b)
1979Parhabdolithusjudithae Black, 1972. Wind& Cepek,p. 231 , pi. 11 ,figs. 5-7.

Derivation of name: Latin. bulla/us = inflated.
Diagnosis: A species of Percivalia with a hollow, bulbous
distal projection that fills the central area.
Description (LM only): Medium-sized, elliptical murolith
exhibiting a bicyclic rim extinction pattern. The narrowouter rim is slightly less birefringent than the inner rim
cycle, which grades into a finely granular central area of
equal, 1st order white birefringence. Extinction lines
extend from the juncture of the two rim cycles into thecentral area. A hollow, bulbous distal projection nearly
fills the central area. The rim elevation is low; the stemheight is about two times that of the rim. Size of observed
specimens: 6.0-8.0 !J.m.
~emar~: Wind & Cepek (1979) illustrated a specimen
from DSDP Hole 397 A as Parhabdoli thus judithae Black,1972. However, the original specimens illustrated fromthe Albian Gault Clay by Black (1972; pl. 3, figs . 5, 6; pl.
4, fig. 4) have high rims and truncated distal projections.

Range: DSDP Site 534 (57% Sa) : late Berriasian toearliest Hauterivian, Cores 83-4, 96-97cm to 70-6, 2730cm (isolated specimen in 67-1, 80-83cm consideredreworked); SE France (Angles, 35% Sa): lo~r Valanginian
to basal Hauterivian, Bed 199 (base otopeta Zone) to Bed
386 (radiatus Zone).
Holotype: Plate 1, figs. 4a-b; Angles, Bed 3.34, upper
Valanginian (verrucosurn Zone).
Paratype: Plate 1, figs . 6a-b; Angles, Bed 352, (trinodosumZone).
Pickelhaube umbellatum, new species(Plate 1, figures 19a-c, 20a-b, 2la-b)

Derivation of name: Latin. umbel/a= para~l, sunshade.
Diagnosis: An irregular-shaped species of Pickelhaubewith two faintly birefringent shields of low element number; the proximal shield elements are strongly curved.

========================~63========================~

Journal ofNannoplankton Research, 16, 2, 1994.

Description: Medium to large placolith constructed of
two faintly birefringent shields. The larger proximal
shield is constructed of 13-19 elements with strongly
curved sutures. The horizontal periphery is highly irregular, although the outline is basically sub-circular to circular.
The small, narrow distal shield is about 1/3-1/4 the size ofthe proximal shield. A brightly birefringent central crossnearly fills the small central area and has arms that taper
to points. A long, proximally-directed stem is present onsome specimens and appears to have octoradiate symmetry. Size of observed specimens: 6 .4-8.8 J..lm.
Remarks: Pickelhaube furtiva (Roth) has brightly
birefringent shields, a smooth horizontal peripheral outline, and radial proximal shield elements.
Range: Rare, extremely short stratigraphic range within
the middle lower Valanginian. DSDP Site 534 : Sample
79-5, 120-123cm; SEFrance(Angles, lOO% Sa): Bed258
to 274 (lower campylotoxum Zone).

J. A. Bergen: Beniassian to Early Aptian Calcareous Namof06Gils. .. , p. ~9 -69.

Upper Cretaceous. It is optically distinct from younger
taxa within this lineage, which have broader rims (with apitted or a serrate inner margin) and/or transverse bars
that exhibit a diagonal, bright birefringence band (whenparallel to the polarizing direction).
Range: Cosmopolitan; mid-Hauterivian to Santonian. Its
LOin the following sections: (1) DSDP Site 534,59-3,7679cm (B 9/10 Sa); (2) SE France (Vergons, B 9/10 Sa),Bed 92, savni Zone; (3) Speeton {approximate), Bed CS.
Holotype: Plate 1, figs . 24a-c; SpeetonBorehole, 11.01m.

Rhagodiscus de!«Jene/ii, new species(Plate 1, figures 7a-c, 8a-d, 9a-b)
1979 Parhabdolithus swinnertonii (Black, 1971) Wind & Cepek, p. 231 ,pi. 11, figs. 8-16.

Derivation of name: named in honour of Eric de Kaenel.
Holotype: Plate 1, figs. 19a-c; DSDP Site 534, Core79-5, 120-123cm.

Reinhardtites scutula, new species(Plate 1, figures 24a-c, 25a-b)
1989 Zeugrhabdotus sisyphus (Gartner, 1968) nov. comb. - Crux, p. 198,
pi. 8.7, fig . I, pi. 8.12, fig. 30.
(?) 1991 Zeugrhabdotus sisyphus (Gartner, 1968)Crux, 1989- Mutterl<'6e,p. 131 , pi. 9, figs. 3, ~ .

Derivation of name: Latin. scutula = diamond or
lozenge-shaped figure.
Diagnosis: A species of Reinhardtites having a smooth,narrow rim and a central, transverse bar supporting a
distal boss of equal width. A bright, elongated, diamondshaped extinction figure is observed on the central portionof the faintly birefringent, transverse bar.

Diagnosis: A species of Rhagodiscus with a brightly
birefringent, solid distal stem; the stem tapers distally andits horizontal periphery extends to, or outside, the central
area margin.
Description: Medium-sized, elliptical murolith. The distal
shield is constructed oflath-shaped, dextrally-imbricatedelements with radial distal sutures. The proximal shield isnarrow and thin. The thick, imperforate central area iscomposed of granular elements. A solid, tapered distal
projection nearly fills the central area and occupies 1/2 to
2/3 of the coccolith width (reaching to or extending
beyond the central margin). Its height is equal to, or isslightly less, than the height of the rim. The stem has aserrate horizontal periphery and is constructed of twoelement cycles. The large, outer cycle is non-imbricate ( 10or more elements); a small central cycle can be observed
on some specimens. The stem is brightly birefringent andis divided into quadrants by the extinction lines. Size ofobserved specimens: 4.8-6.4 J..lm.

Description: Medium to large, normally to narrowly
elliptical murolith. The distal shield is constructed of asingle cycle of 32-40 elements with dextral imbrication.
The central area is greater than 1/2 the coccolith width(0.53 to 0.67 relative width). An elevated, transverse barspans the central area and is constructed of numerous
fibrous elements oriented parallel the minor ellipse axis.
A circular distal boss equal in width to the transverse bar
is present at its centre. The rim extinction pattern isunicyclic and displays a first order grey birefringence;extinction lines spiral across the rim. The transverse bar
exhibits same faint birefringence as the rim; when specimens are oriented 45 degrees to the polarizing direction,
the bar is divided by a longitudinal extinction line. A
brightly birefringent, elongate, diamond-shaped figure isobserved on the central portion of the bar when specimens
are oriented parallel to the polarizing direction. Measured
specimens ranged from 6.5-8.5 J..lm in length and 1.3-1.6
in ellipticity.

Range: early Valanginian to early Hauterivian. DSDP
Site 534 (100% Sa): 80-1, 0-3cm to 63-2, 66-69cm. SE
France (90% Sa): base camovlotoxum Zone (Angles, Bed250) to base nodosoplicatum Zone (Vergons, Bed 57).
Also reported from DSDP Site 397 A (Wind & Cepek,1979) and Site 603 (Covington &.:Wise, 1987), ODP Sites
638-641 (Applegate & Bergen, 1988).

Remarks: Reinhardtites scutula is the ancestral species
ofa lineage that has excellent biostratigraphic u~lity in the

Holotype: Plate 1, figures Sa-d, DSDP Site 534,71-1,6366cm, upper Valanginian.

Remarks: The holotype of Rhagodiscus iwinnertonii(Black) Applegate et al., 1987 was recovered from theAptian Sutterby Mar!; its stem does not extend to thecentral area margin and is constructed of numerous
elements showing strong laevogyre curvature. Both theappearance and extinction of Rhagodiscus de!«Jenelii areexcellent biostratigraphic datums in low.:latitude anddeep-sea sections.

========================~64=========================

J. A. Bergen: Berriassian to Early~ Calcareous Nannofossik .. , p. 59-69.

Rucinolithus pinnatus, new species(Plate l, figures 3a-b)
1989 Rucinolithus terebrodentarius Applegate et al., 1987. Bralower,
Monechi, & Thierstein (partirn), p. 223, pi. 7, figs. 4-6 (non pi. 7, figs.

1-3).

Derivation of name: Latin. pinnatus = feathered, plumed.
Diagnosis: A species of Rucinolithus possessing eightelements that broaden outwards along the sutures between adjoining elements and then taper to points once theelements become separate.
Description: Medium-sized, robustnannolithconstructedof eight, imbricated elements with curved sutures. The
elements are broadest at their outer peripheral contact
and then taper outwards to form pointed terminations. A
small element cycle surrounds a small central perforation
or boss. Size of holotype: 6.9J.lm.
Remarks: Bralower et al. (1989) reported the range of
Rucinolithus terebrodentarius as upper Hauterivian toTuronian, but one of their illustrated specimens wasrecovered from Berrasian section (other illustrated specimens from the same sample include Diadorhombus
rectus, Umbria granulosa ssp. granulosa, andRhagodiscus
nebulosus) and serves as the holotype for Rucinolithus
pinnatus. Nannoconus ligius is another robust Neocomiannannolith with a petaloid form and eight-fold symmetry.
It is possible that Rucinolithus pinnatus is an isolated
distal calyx; this was demonstrated for another nannolithwith eight-fold symmetry, Retecapsa radiata (Worsley)
Applegate & Bergen. R. pinnatus is distinguished from
Kokia spp. by the strong imbrication of the elements.
Range: Very rare, composite range ofmid-Tithonian tolatest Berriasian. DSDP Site 534 (34% samples): Cores
96-3 , 9-12cm to83-l, 18-19cm; SE.France, Angles (55%Sa), upper Berriasian, Bed 158 to Bed 180.
Holotype: Bralower, Monechi, & Thierstein ( 1989),
plate 7, figures 4-6; DSDP Site 100, 1-1, 13cm, Berriasian.

Vagalapilla, species A(Plate 1, figures 28a-b)

Journal ofNannoplankton Research, 16, 2, 1994.

GENUS Zeugrhabdotus Reinhardt, 1965
Remarks: The large number of Cretaceous muroliths
having transverse central bars show a great range in opticalproperties and ultrastructure. Four species cited and/or
described herein have been retained within the Mesozoic
genus Zeugrhabdotus, whereas four others are left within
the Palaeogene genus Zygodiscus (see list under Other
Species Considered). The proper classification of suchforms cannot be made until the ultrastructure and evolutionof this entire murolith family is better understood, especially their transitions across the Cretaceous/Tertiaryboundary.

Zeugrhabdotus trivectis, new species(Plate I , figures 26a-b, 27a-c)
Derivation of name: Latin. trinus = three. vectis = bar,lever.
Diagnosis: A species of Zeugrhabdotus exhibiting a bicyclic rim extinction pattern of contrasting birefringence(inner cycle is bright); optically, the transverse bar gives
the impression ofthree element bundles arranged at slightly
oblique angles to each other.
De~cription: Medium-sized, normally elliptical murolith.
The distal shield is constructed of a single cycle of 28-3 2
elements with strong dextral imbrication. A thin, narrow,basal inner cycle observed in distal view may represent the
inner portion of the proximal shield. The central area isspanned by a transverse bar constructed of fibrous elements. The bar broadens slightly near the juncture with theinner rim margin. A circular distal projection, which isnarrower than the bar, is present at the centre of thecoccolith. Specimens exhibit a bicyclic rim extinction
pattern. The inner cycle displays a 1st order whitebirefringence and the outer cycle is faintly birefringent.
The transverse bar displays the same bright birefringenceas the inner rim cycle when specimens are oriented parallelto the polarizing direction. The bar elements are opticallygrouped into three bundles oriented at slightly obliqueangles to each other. Measured specimens: length 4.7-6.0
J.lm; ellipticity 1.3-1. 5; relative central area size 0.50-0.57.

Remarks: As with most Zeugrhabdotus species, the optical properties of Zeugrhabdotus trivectis are more distinct
than its ultrastructure.

1966 Coccolithus matalosus Stover, p. 139, pi. 2, fig. 2 (non pi. 2, fig. I,
pi. 8, fig. I 0).

Remarks: TheholotypeofVaga/api//a matalosa (Stover,1966) Thierstein, 1973 (see Stover, 1966, pl. 2, fig. 1) isa placolith belonging to the genus Broinsonia Bukry,
1969; its lowest occurrence in the Vocontian Troughsections is lowermost Albian. The other photographed
specimen in Stover ( 1966) is a murolith, which has adiamond-shaped central opening. Very rare specimens
having elliptical central openings were recovered fromthe upper Hauterivian.
Range: early Aptian to early Cenomanian. SE France:
LO at La Bedoule (92% Sa), Bed 86.

Range: Cosmopolitan, early Valanginian to middle
Cenomanian. LOin the following sections: (1) DSDP Site
534 (28%Sa), Core77-3, 43-44cm; (2) SEFrance(Angles,
70% Sa), Bed 284, campylotoxum Zone.
Holotype: Plate 1, figure 27a-c, BGS Speeton Borehole,
14.07m.

Zeugrhabdotus species A(Plate 1, figure 16a-b)
Description (LM only): Medium-sized, normally elliptical murolith. The rim exhibits a uni.~orm 1st order grey
birefringence. An oblong, transverse bar spans the central
area and has pointed terminations that extend into the rim.

========================~65===========================

J. A. Bergen: Beni~ian to Early Ap.ian Calcareous Nannofossils... , p. 59- 69.

Journal ofNamoplankton Research , 16, 2, 1994.

The transverse bar displays a l st order white birefringencethat strongly contrasts with the birefringence of the rim.
A central, circular distal projection is. non-birefringent,interrupting the continuity of the birefringent transverse
bar. Longitudinal extinction lines further subdivide thetransverse bar. Size of observed specimens 4 .8-5 .8 Jlm.
SPECIESC. oblongata
C. tenuis
C. mexicana
C. geometricum
C. silvaradion
C. delicatus
C. cuvillieri
C. brezae
C. deflandrei
D . rectus
D . primitiva
E. primus
E. striatus
E. windii
E. antiquus
E. floral is
E. hauterivianus
F. multicolumnalus
F. oblongus
H. circumradiatus
H . ellipticus
H . irregularis

DSDP 534 {Core-Section, cm)
Freq.
LO
HO
95%
80-5, 121
50-3, 7168%
76-6,94---------

--T9
B3!f9
87
95%
80%
T9
84
24%

T2
43%
77%
1Sa

---T5
62%
1Sa
100%
67%

---

-----------------

82-1. 132
86-3,40
91-2, 145
95-3,64-·-------85-3 , 108
67-1, 80--------67-1, 80
79-5, 120-------56-1,49
59-3,76--------59-3,76
63-2,66
78-2,85--------51-1, 61
78-2,85
63-2,66
70-2,50

64-4,25----------------106-1, 54
44-4, 1186-3,40104-1---------

64-4,25--------50-3,70
92-1, 79--------49-2,70------------------

H. noelae
L . bollii--64%96-3,966-3,60

91-2, 145
52-3,72

L.carniolensis
M . mecurius
M . speetonensis
M . conus
N . compressus
N . ligius
N . steiNnannii
P . fenestrata
P . senmia
R . angustiforata
R. radiata
R. achlyostaurion
R. angustus
R. nebulosus
R. wiseiS . lowei
S . colligata
S. mutterlosei
S . comptus
T. gabalus
T. jurapelagicus
T. verenae
U. granulosa
V. mitcheneri
V. parallela
Z. erectus
Z.[ISSUS
Z. bicrescenticus• Z. diplogrommv.s
Z. elegans
Z. xenotus

92%
1Sa .

88-4, 14176-2, 61--------76-2, 61--------75-2,49
89-1,26-----------------------

70%

85-4, 126
97-1, 1956-1,49
91 -2,45
83-4,96
93-2,4191 -1, 80
69-2,95
45-4, 10--------88-2, 17
83-1, 18--75%--100%
57%
96%
33%----100%
810
--94%
43%
1Sa
16%
89
89

TlO
57%
83
T8
T8
85
B8

--86

--------91 -1, 80
78-2,85
71 -1, 63
80-5, 12985-4, 126
83-2,92
96-3, 968-5, 1
79-2, 10--------116-1
70-2,50
78-3,49
48-2, 110
77-3, 111-·-----------------89-4,35---------47-2,80-----·-------------82-2,49
76-2, 61
52-3,72
54-2,57-- -·- -·--·-71-1, 63-----------------67-1, 80
83-2,92
49-3,60--------53-2,60
70-6,27----------------------·------------

Remarks: Specimens must be transferred from the LM to
the SEM before this taxon can be recognized either as avalid or new species. It is closely related to Zeugrhabdotus
erectus, from which it is distinguished by the opticalproperties of its central structure (see Plate I) .
SE France (Section, Bed I#)
Freq.
LO
Angles 1, 199
99%
74%
Angles 1, 275
80%---------·-------absent
B3{f2Angles 1, 170
Angles, 126
308100%--------------·B8(f4Route de Grads, 1
T5-----------·---·86
Angles 1, 164
Angles 1, 299
2Sa
T8---------------Angles 1, 354
98%
96%
Angles 1, 258
1Sa
100%

T2

---

43%
98%

--87

--95%

--60%
80
2Sa

--89

--100%

Vergons, 51
La Bedoule, 169-------·------bimammaJum Zone
Angles 2, 147
Angles 1, 102
Route de Grads, 25
Angles 2, 199
La Bedoule, 59
fallaJ,Ui Zone
Vergons,48
Angles 1, 77 (?)
Angles 1, 305
Angles 1, 250
Angles 1, 258
fallaJ,Ui Zone
Vergons, 141
Angles 1, 79
Angles 1, 132--------------B8!f9
B2!f9
85
85%

--B8(f3
---

TlO
13%
2Sa
9%
88
B6(f6
T7
19
85
T5
T6
86
89
--87

Angles, 79
Vergons,40
Angles 2, 165
la Bedoule, 129
Route de Grads, 27?
Angles 1, 167---------------Route de Grads, 27
Angles 1, 321
Angles 1, 306b
Angles 1, 180
Angles 1, 131a
Angles 1, 139---------------Vergons,32
Angles 1, 264--------------la Melle, Oxfordian
Angles 1, 311
Angles 1, 258
Angles, u. Aptian
Angles 1. 278

HO
Angles 2. 116--------La Bedoule, 114
absentVergons, 110-----·- --Vergons, 110
Vergons, 68
Angles 1, 278-------·Angles 1, 306b
Angles 1, 278
Vergons, 78
Angles 1, 397;
Vergons, 19
Vergons, 51--------------Angles 2, 94
Route de Grads, 25------------------------------

Angl~s 2. 90

Blieux, u. Albian
Route de Grads, 25
Vergons, 196
Angles 2. 52--------------Angles 328
Angles 2. 261
Angles 1, 264
Route de Grads, 27
Angles, u. Aptian
La Bedoule, 161---·-----------Route de Grads, 27
La Bedoule, 114
Angles 2. 134-----------------·----------·--

Angles 1, 158 ·Angles 1, 306a
Angles 2. 79
Vergons, 141
lower CenomanianAngles 1, 309-·--------------lower Turonian
Vergons, 96
Angles 1, 147
Vergons, 123
Blieux, Albian
Vergons, 160
Angles 1, 324-------------·----------------~i&ii~-c;~manian

Table 1: Biostratigraphic Summary (excluding species described or recombined here). Freq. refer.; to either: (I) the percentage of samples containing
species within its stratigraphic range (e.g. 70%); or(2) the number of samples in which a species was observed either in the top (T) or bottom (B) ten samples
of its strati graphic range (e.g . T3 means specimens observed in 3 of I 0 samples at the to'p of its strati graphic range). Angles I refer.; to the BerriasianValanginian section and Angles 2 to the Barremian stratotype along Routed ' Angles.

===========================66===========================

J. A. Bergen: Beniassian to Early Aptian Calcareous Nannofa.rus. .. , p. 59- 69.

Journal ofNannoplankton Research, 16,2, 1994.

Range: latest Hauterivian to early Campanian. Its LO in
the following sections: (l)DSDP Site534, 52-1, 82-85cm;(2) SE France, Angles, Barrernian stratotype, Bed 60,
angulicostata Zone; (3) Speeton (approximate), upper
Hauterivian, Bed C2, rnarginatus Zone.

Vekshinella para/le/a (Wind & Cepek, 1979) Applegate& Bergen, 1988.
Zeugrhabdotus erectus (Deflandre, 1954) Manivit, 1971.
Zeugrhabdotusfissus Griin & Zweili, 1980.
Zygodiscus bicrescenticus (Stover, 1966) Wind & Wise;
1976.
Zygodiscus diplogrammus (Deflandre, 1954) Gartner,
1968.
Zygodiscus elegans Gartner, 1968.
Zygodiscus xenotus (Stover, 1966) Hill, 1976.

Otber Species Considered (by Generic Epithet)
Assipetra infracretacea (Thierstein, 1971) Roth, 1973.
Calcicalathina oblongata (Worsley, 1971) Thierstein,
1971.
Chiastozygus tenuis Black, 1971.
Corollithion geometricum (G6rka, 1957) Manivit, 1971 .
Corollithion si/varadion (Filewicz et al., 1987) PerchNielsen, 1984.
Conusphaera mexicana Trejo, 1969.
Cretarhabdus delicatus Applegate, Covington, and Wise,
1987.
Cruciellipsis cuvillieri (Manivit, 1966) Thierstein, 1971.
Cyclagelosphaera brezae Applegate & Bergen, 1988.
Cyclagelosphaera dejlandrei Manivit, 1966.
Diadorhombus rectus Worsley, 1971.
Diloma primitiva (Worsley, 1971) Wind & Cepek, 1979.
Eiffellithus primus Applegate & Bergen, 1988.
Eiffel/ithus stria/us (Black, 1971) Applegate & Bergen,
1988.
Eiffel/ithus windii Applegate & Bergen, 1988.
Eprolithus antiquus Perch-Nielsen, 1979.
Eprolithus flora/is (Stradner, 1963) Stover, 1966.
Ethmorhabdus hauterivianus (Black, 1971) Applegate etal., 1987 in Covington & Wise, 1987.
Faviconus multicolumnatus Bralower, 1989.
Flabellites oblongus (Bukry, 1969) Crux, 1982.
Haqius circumradiatus (Stover, 1966) Roth, 1978.
Haqius el/ipticus (Griin, 1975) Bown, 1992.
Hayesites irregularis (Thierstein, 1971) Applegate et al.,1987 in Covington & Wise, 1987.
·
Hexa/ithus noelae Loeblich & Tappan, 1966.
Lithraphidites bol/ii (Thierstein, 1971) Thierstein, 1973.
Lithraphidites carniolensis Deflandre, 1963.
Metadoga mecurius Wind & Cepek, 1979.
Micrantholithus speetonensis Perch-Nielsen, 1979.
Microstaurus conus Wind & Cepek, 1979.
Nannoconus compressus Bralower & Thierstein, 1989.
Nannoconus ligius Applegate & Bergen, 1988.
Nannoconus steinmannii Karnptner, 1931
Percivaliafenestrata (Wors1ey, 1971) Wise, 1983.
Po/ycostel/a senaria Thierstein, 1971.
Retecapsa angustiforata Black, 1971 .
Retecapsa radiata (Worsley, 1971) Applegate & Bergen,
1988.
Rhagodiscus achlyostaurion (Hill, 1976) Doeven 1983.
Rhagodiscus angustus (Stradner, 1963) Reinhardt, 1971 .
Rhagodiscus nebulosus Bralower, 1989.
Rucinolithus wisei Thierstein, 1971 .
Sollasites horticus (Stradner et al., 1966) Cepek & Hay,
1969.
Sol/asites lowei (Bukry, 1969) Roth, 1970.
Speetonia col/igata Black, 1971.
Staurolithites mutter/osei Crux, 1989.
Stradnerlithus comptus Black, 1971.
Tranolithus gabalus Stover, 1966.
Tubodiscusjurape/agicus (Worsley, 1971) Roth, 1973.
Tubodiscus verenae Thierstein, 1973.
Umbria granulosa Bralower & Thierstein, 1989.
Vekshinella mitcheneri Applegate & Bergen, 1988.

REFERENCESApplegate, J.L. and Bergen, J.A. , 1988: Cretaceous calcareous
nannofossil biostratigraphy ofsediments recovered fromODP Leg 103, Galicia Bank. IRDSDP, 3, 293-348.
Bralower, T.J., Monechi, S., and Thierstein, H.R., 1989: Calcareous nannofossil zonation of the Jurassic-Cretaceous
boundary interval and correlation with the Geomagnetic
Polarity Timescale. Mar. Micropa/eont., 14, 153-235.
Busnardo, R. , 1984 : CrcHace inferieur: echelles
biostratigraphiques. In: " Synthese geologique du SudEst de la France", Mem. Bur. Rech. geo/. Min ., 125,291-294.
.
Busnardo, R. , Atrops, F., Crumiere, J.-P., Dromart, G.: FieldTrip Amoco, South-east of France, Guide Book, May
10-25, 1987.
Covington, M. & Wise, S.W., 1987: Calcareous nannofossil
biostratigraphy of a Lower Cretaceous deep sea fancomplex: DSDP Leg 93, Site 603,lowercontinental riseofT Cape Hatteras. IRDSDP, 93,617-660.
Crux, J.A., 1989: Biostratigraphy and paleogeographical applications of Lower Cretaceous nannofossils from
north-western Europe. In, Crux, J.A. & van Heck, S.E.
(eds.), Nannofossi1s and their Applications, British
Micropalaeontol. Soc. Ser., 143-211.
Hoedemaeker, P.J. and Bulot, L.G. (reporters), 1990: Preliminary ammonite zonation for the Lower Cretaceous ofthe
Mediterranean region: report. Geologie alp, 66, 123-127.
Hoedernaeker, P.J. and Company, M. (reporters), 1993: Ammonite zonation for the Lower Cretaceous of theMediterranean region; basis for the stratigraphic correlation within I. G.C .P. Project 262 . Revta. Esp. Palaeont.,
8, 117-120.
Roth, P.R., 1983: Jurassic and Lower Cretaceous nannofossils
in the western North Atlantic (Site 534): biostratigraphy, preservation, and some observations on biogeography and paleoceanography. IRDSDP, 76,731-759. .
Sissingh, W., 1977: Biostratigraphy of Cretaceous calcareous
nannoplankton. Geologie Mifnb., 56, 37-65.
Sissingh, W., 1978 : Microfossil biostratigraphy andstage-stratotypes of the Cretaceous. Geo/ogie Mijnb.,
57, 433-440.
Thierstein, H:R., 1971 : Tentative Lower Cretaceous calcareous
nannoplankton zonation. Ec/og. geo/. Helv., 64,459-488.
Thierstein, H.R., 1973 : Lower Cretaceous calcareous
nannoplankton biostratigraphy. Abh. geol. Bundesanst.,
29, 1-52.
Wind, F.H. and Cepek, P., 1979: Lower Cretaceous calcareous
nannoplankton from DSDP Hole 397A (northwest
African Margin). IRDSDP, 4711, 221-255 .
APPENDIX;.~~
D.S.D.P. Site 534 samples examined: 44-4, ll-14cm; 45-4,
10-13cm; 46-1, 99-102cm; 47-2, 80-83cm; 48-2, 110-113cm;
49-2, 70-73cm; 49-3,60-63cm; 49-4,61-64cm; 50-I, 68-71cm;
50-3, 71-74cm; 51-1,61-64cm; 52-1, 82-85cm; 52-3, 72-75cm;======================~67======================~

Journal ofNannoplankton Research, 16, 2, 1994.

J. A. Bergen: Beni~an to Early AJXian Calcareous Nannof<l&'>ils. .., p. 59- 69.

53-2, 60-63cm; 54-2, 57-60cm; 55-3, 75-78cm; 56-1, 49-52cm;
58-1, 42-45cm; 59-3, 76-79cm; 60-3, 52-55cm; 61 -2, 80-83cm;
63-2, 66-69cm; 64-2, 78-81cm; 64-4, 25-28cm; 65-3, 102195cm; 65-6, 69-72cm; 66-3 , 60-63cm; 67-1, 80-83cm; 68-5,
1-4cm; 69-2, 95-98cm; 70-2, 50-53cm; 70-6, 27-30cm; 71-1 ,63-66cm; 72-4, 12-15cm; 73-1 , 125-1 28cm; 74-6, 73-76cm; 752, 49-52cm; 75-6, 103-107cm; 76-2, 61-64cm; 76-6, 94-95cm;
77-3, 43-44cm; 77-3 , 111-114cm; 78-2, 85-86cm; 78-3, 4952cm; 78-5 , 7-8cm; 79-2 , 10-11cm; 79-5, 120-123cm; 80-1 ,0-3cm; 80-2 , 80-82cm; 80-5 , 129-130cm; 81-3, 57-58cm; 81-4,
48-51cm; 82-1 , 132-133cm; 82-2, 49-52cm; 83-1 , 16-18cm; 832, 92-95cm; 83-4 , 96-97cm; 84-1 , 89-90cm; 84-2 , 129-132cm;

84-3, 113-114cm; 84-4, 41-42cm; 84-5, 86-87cm; 85-2, 5154cm; 85-3, 108-109cm; 85-4, 126-127cm; 86-1, 122-125cm;
86-3, 40-41cm; 86-5, 36-37cm; 87-2, 64-67cm; 87-4, 40-41cm;
87-5, 122-123cm; 88-2, 17-20cm; 88-4, 141-142cm; 89-1 , 2627cm; 89-4, 35-36cm; 90-1, 123-124cm; 90-3, 21-23cm; 91-1,
80-81cm; 91-2,45-46cm; 91-4, 131-132cm; 91-5, 134.-135cm;
91-6,1 -2cm; 92-1, 79-80cm; 92-2, 131-132cm; 92-3, 109-110;
92-4, 147-148cm; 92-5, 69-72cm; 92-5, 86-87; 92-6, 48-49cm;
93-1 , 105-106cm; 93-2,41-44cm, 93-2,100-103cm; 93-3,122125cm; 94-1, 93-94cm; 94-2, 89-92cm; 94-3, 89-92cm; 94-4,
3-4cm.

PLATE 1
All LM 2500X. XP - cross ·polarized light; Ph - Phase contrast illumination; Tr - transmitted light.

Hexalithus strictus, n. sp.; (1) holotype, DSDP 534, 91-4, 131-132cm, (a) XP, (b) Tr; (2) DSDP 534, 89-4, 3536cm, (a) XP, (b) Tr.
Rucinolithus pinnatus, n. sp., Angles, Bed 164, (a) XP, (b) Tr.
3:
Percivalia bullata, n. sp., Angles, (4) holotype, Bed 334, verrucosum Zone, (a) XP, (b) Tr; (5) Bed 364,
4-6:trinodosum Zone, (a) XP, (b) Tr; (6) paratype, lateral view, Bed 352, trinodosum Zone, (a) XP, (b) Tr.
7-9:
Rhagodiscusdekaenelii, n. sp.; (7) DSDP Site 534, 71-1 , 63-66cm, u. Valanginian, (a)XP, (b) Tr, (c) proximal
view; (8) holotype, DSDP Site 534, 71-1 , 63-66cm, u. Valanginian, (a) lateral view, (b) distal view, (c) XP,(d) Ph; (9) Angles, Bed 357, trinodosum Zone, (a) XP, (b) Tr.
10:
Grantarhabdus quadratus (Worsley) n. comb., Speeton, Bed C11A, regale Zone.
Ca/cica/athina sp. A, Angles, Bed 245, u. Berriasian, (a) XP, (b) Tr.
11:
Diloma sp. A, Speeton, Bed LB5, lo. Barremian, (a) XP, (b) Tr.
12:
13-14: Diloma galiciense, n. sp. (13) DSDP 534, 59-3 , 76-79cm, u. Hauterivian, (a) XP, 45°, (b) XP, 0°, (c) Tr; (14)DSDP 534, 52-l , 82-85cm, u. Hauterivian, (a) XP, 45°, (b) XP, 0°, (c) Tr.
Zeugrhabdotus erectus (Deflandre), Speeton, Bed Cl lA, regale Zone, (a) XP, (b) Tr.
15:
Zeugrhabdotus sp. A, Angles, upper Aptian, (a) XP, (b) Tr.
16:
17-18: Lucianorhabdus salomonii, n. sp. ( 17) holotype, lateral view, Angles, Barremian, Bed 161, sartousiana Zone,(a) XP, (b) Tr; (18) plan view, rim, La Bedoule, Bed 157, (a) XP, (b) Tr.
19-21: Pickelhaube umbellatum, n. sp., lo. Valanginian; (19) holotype, DSDP 79-5, 120-123cm, (a) lateral view, (b)XP, (c) Tr; (20) Angles, Bed 274, campylotoxum Zone, (a) XP, (b) Tr; (21) DSDP 534,79-5, 120-123cm, (a)
1-2:

~Mh

.

22-23: Markalius vetulus, n. sp., Angles, campylotoxum Zone; (22) holotype, Bed 278, (a) XP, (b) Tr; (23) Bed 302,(a) XP, (b) Tr.
24-25: Reinhardtites scutula, n. sp. (24) holotype, Speeton borehole, 11.01m, (a) distal view, (b) XP, (c) Tr ; (25)
Speeton, Bed LB5, lo. Barremian, (a) XP, (b) Tr.
26-27: Zeugrhabdotus trivectis, n. sp. (26) Angles,Bed 361 , trinodosum Zone, (a) XP, (b) Tr; (27) holotype, Speeton
borehole, 14.07m, (a) distal view, (b) XP, (c) Tr.
Vagalapilla, sp. A, La Bedoule, Bed 153, (a) XP, (b) Ph.
28:===========================68===========================

Journal of annoplankton Resear h, 16, 2, 1994.

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